Mammals - The Zambezi Society

Transcription

AWF FOUR CORNERS TBNRM PROJECT : REVIEWS OF EXISTING BIODIVERSITY INFORMATION
i
Published for
The African Wildlife Foundation's
FOUR CORNERS TBNRM PROJECT
by
THE ZAMBEZI SOCIETY and
THE BIODIVERSITY FOUNDATION FOR AFRICA
2004
PARTNERS
IN
BIODIVERSITY
The Zambezi Society
P O Box HG774
Highlands
Harare
Zimbabwe
Tel: +263 4 747002-5
E-mail: [email protected]
Website : www.zamsoc.org
The Biodiversity Foundation for Africa
P O Box FM730
Famona
Bulawayo
Zimbabwe
E-mail: [email protected]
Website: www.biodiversityfoundation.org
The Zambezi Society and The Biodiversity Foundation for Africa are working as partners within the
African Wildlife Foundation's Four Corners TBNRM project. The Biodiversity Foundation for Africa is
responsible for acquiring technical information on the biodiversity of the project area. The Zambezi
Society will be interpreting this information into user-friendly formats for stakeholders in the Four
Corners area, and then disseminating it to these stakeholders.
THE BIODIVERSITY FOUNDATION FOR AFRICA (BFA is a non-profit making Trust, formed in
Bulawayo in 1992 by a group of concerned scientists and environmentalists. Individual BFA members
have expertise in biological groups including plants, vegetation, mammals, birds, reptiles, fish, insects,
aquatic invertebrates and ecosystems. The major objective of the BFA is to undertake biological research
into the biodiversity of sub-Saharan Africa, and to make the resulting information more accessible.
Towards this end it provides technical, ecological and biosystematic expertise.
THE ZAMBEZI SOCIETY was established in 1982. Its goals include the conservation of biological
diversity and wilderness in the Zambezi Basin through the application of sustainable, scientifically sound
natural resource management strategies. Through its skills and experience in advocacy and information
dissemination, it interprets biodiversity information collected by specialists like the Biodiversity
Foundation for Africa and uses it to provide a technically sound basis for the implementation of
conservation projects within the Zambezi Basin.
THE PARTNERSHIP between these two agencies was formed in 1996 as a result of mutual recognition
of their complementarity. They have previously worked together on several major projects, including the
biodiversity component of IUCN's Zambezi Basin Wetland project and the evaluation of biodiversity in
Tete province described in detail in the first Four Corners TBNRM Biodiversity Information Package.
ISBN 0-7974-2835-6
RECOMMENDED CITATION:
Timberlake, J.R. & Childes, S.L. 2004. Biodiversity of the Four Corners Area: Technical Reviews Volume Two
(Chapters 5-15).
Occasional Publications in Biodiversity No 15,
Biodiversity Foundation for Africa, Bulawayo/Zambezi Society, Harare, Zimbabwe.
THE AWF FOUR CORNERS TBNRM PROJECT IS FUNDED BY USAID THROUGH THE REGIONAL CENTER FOR SOUTHERN AFRICA
ACKNOWLEDGEMENTS
In addition to all the authors of the chapters in this document,
The Zambezi Society and the Biodiversity Foundation for Africa
wish to thank the following for peer review or additional assistance:John Burrows
Colin Craig
Barbara Curtis
Cornell Dudley
Kevin Dunham
Peter Frost
Debbie Gibson
Mike Griffin
Michael Irwin
Bas Jongeling
Linley Lister
Gillian Maggs
Anthony Mapaura
Brian Marshall
Susan Ringrose
We also wish to thank Henry Mwima, Nesbert Samu,
Othusitse Lekoko, Daudi Sumba, Gitonga Kathurima,
Maureen Mashingaidze and Simon Metcalfe
of the African Wildlife Foundation
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CHAPTER 6. MAMMAL FAUNA OF THE FOUR CORNERS AREA
CONTENTS
PUBLISHERS’ DETAILS …………………………………………………………………….. i
ACKNOWLEDGEMENTS ..........................................................................................................ii
FIGURES ......................................................................................................................................iv
TABLES ........................................................................................................................................iv
APPENDICES ..............................................................................................................................iv
CHAPTER 6. MAMMAL FAUNA OF THE FOUR CORNERS AREA ...........................231
6.1 INTRODUCTION .........................................................................................................231
6.1.1 Geographical and Taxonomic Scope.......................................................................231
6.1.2 Taxonomic Issues and Uncertainties.......................................................................232
6.2 SOURCES OF KNOWLEDGE .....................................................................................232
6.2.1 Historical Aspects....................................................................................................232
6.2.2 Taxonomic Diversity and Surveys ..........................................................................233
6.2.3 Ecological Studies ...................................................................................................235
6.3 PRINCIPAL ASSEMBLAGES OF MAMMALS.........................................................236
6.3.1 Kalahari Savanna.....................................................................................................236
6.3.2 Mopane Savannas and Dry Miombo Woodlands....................................................238
6.3.3 Mesic Miombo Woodlands .....................................................................................238
6.3.4 Wetlands, Grasslands and Riparian Woodlands......................................................238
6.3.5 Rocky (Rupicolous) Habitats ..................................................................................238
6.4 ECOLOGICAL AND EVOLUTIONARY PATTERNS AND DETERMINANTS.....239
6.4.1 Ecological Determinants .........................................................................................239
6.4.2 Mammals as Ecological Determinants ....................................................................239
6.4.3 Landscape Evolution ...............................................................................................239
6.5 NOTABLE POPULATIONS OF LARGE MAMMALS, INCLUDING HISTORICAL
EXTINCTIONS AND RANGE CONTRACTIONS...............................................................242
6.5.1 Carnivores ...............................................................................................................242
6.5.2 Ungulates.................................................................................................................242
6.6 WATER AVAILABILITY AND MOVEMENTS ........................................................247
6.7 CONSERVATION STATUS AND CONCERNS ........................................................248
6.7.1 Threatened Species and Habitats.............................................................................248
6.7.2 Diseases ...................................................................................................................248
6.7.3 Introductions of Threatened And Extinct Species...................................................249
6.8 MONITORING AND FUTURE STUDIES ..................................................................249
6.8.1 Monitoring Large Mammals of High Conservation Concern .................................249
6.8.2 Ecological Studies ...................................................................................................250
6.8.3 Biodiversity Surveys, Monitoring and Taxonomic Research .................................250
6.9 MAMMAL CONSERVATION VALUES OF THE FOUR CORNERS TFCA...........251
6.10
REFERENCES ...........................................................................................................252
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FIGURES
Figure 6.1. Distribution of collecting sites of all known bat specimens (Chiroptera) from Angola
showing an acute gap in knowledge, especially in the east of the country. Each marked quarterdegree square represents at least one specimen. After Crawford-Cabral 1989. ......................... 233
Figure 6.2. Extent of alluvial deposits in south-central Africa. Data collated from geological
survey maps for Katanga, DRC and Zambia; extent of alluvium in NE Botswana modified from
Moore & Larkin (2001). Adapted from Cotterill (2003e) and Broadley & Cotterill (2003)...... 240
Figure 6.3. Distributions of the six subspecies of Burchell's zebra. The thick dotted line
approximates the southern limit to modern distribution of zebra in southern Africa, where Equus
b. burchelli is extinct. Modified from Grubb (1981). ................................................................. 244
TABLES
Table 6.1. Mammal species that show an association with the principal faunal elements in the
Four Corners area........................................................................................................................ 237
APPENDICES
Appendix 6.1. Mammals occurring in the Four Corners area, by country. ................................ 260
Chapter 6: Mammals of the Four Corners Area
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F.P.D. (Woody) Cotterill
Lycaon pictus,Wild dog
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F.P.D. (Woody) Cotterill
6.1
INTRODUCTION
The review given in this chapter covers all extant and historically extinct mammals occurring in
the Four Corners Transfrontier area. When its historical context is considered, our current
knowledge of this fauna has grown through surveys and ecological studies of selected species
and assemblages that were carried out independently and piecemeal. Ecological studies have
been biased toward the large mammals (especially in protected areas). Reference to all these
studies has been made, but some information has undoubtedly been missed, especially that in
unpublished reports. It is impossible at present to quantify the quality and scope of this grey
literature.
The principal influences of habitat and faunal associations on the mammal composition are
discussed. In addition, important influences on the evolution of this fauna are singled out and
discussed. Knowledge is summarised for particular species considered to be of conservation,
ecological or economic importance. First, though, the geographical and taxonomic scope of the
mammals is discussed, with brief consideration of the criteria for determining the occurrence of a
species in the region.
6.1.1
Geographical and Taxonomic Scope
Determination of the occurrence of a species within the Four Corners area (Appendix 6.1) has
been derived from the principal summaries of knowledge. With the exception of large, obvious
mammals, the criterion for inclusion has been a reliably identified museum specimen. The list is
thus conservative as it excludes, for example, approximately five species of Chiroptera (bats)
known from nearby localities, but is thought to be representative. Furthermore, many small
mammals as currently listed are likely to comprise two or more species. Important records
omitted include those from the Barotse floodplain and its surrounding miombo woodland (e.g.
Cotterill 2000a), including species of small mammals which could well occur in the Zambian
part of the area. There are an indeterminable number of species of small mammals (especially
shrews and bats) the occurrence of which is based on very sparse data. Examples include
Laephotis botswanae, Kerivoula lanosa, Nycteris macrotis, Mimetillus moloneyi, Scotoecus
hindei and S. albigula, all of which are known from either a single specimen or occur near the
northern boundary. Domesticated mammals are not considered.
The mammal fauna comprises at least 197 species, the majority of which (145 species, or 74%)
are small-bodied (less than 10 kg in mass). The largest groups are rodents (47 species), bats (53
species) and insectivores (9 shrews and one hedgehog), which comprise 56% of the total for the
area.
Besides the great disparity in geographical size of countries, factors such as habitat diversity,
biogeographic affinities and evolutionary history influence these totals profoundly. Not least, the
figures show important limitations in our knowledge of mammals (and in fact all biodiversity) in
each of the five countries. The total species richness of mammals in the area probably exceeds
200, and species are likely to be more widespread than indicated in Appendix 6.1, especially for
Angola.
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6.1.2
Taxonomic Issues and Uncertainties
The taxonomic affinities of the majority of small and many large mammals are very uncertain.
This acute deficiency in knowledge is surprising to many people, including the majority of
biologists. Problems associated with imprecise taxonomic information are serious for both
conservation and wildlife management - the impacts of such inaccurate information on reliable
decisions about populations or habitats should not be underestimated.
Their taxonomy, in terms of what exactly what their constituent species are, is unresolved for
many small carnivores and many genera of elephant shrews, bats, rodents and shrews. Others for
which taxonomy awaits elucidation include primates, notably the baboons and vervet monkeys.
For example, it is impossible at present to state with any confidence what actual species
comprise the rodent genera Cryptomys, Graphiurus and Mus, or the bat genus Scotophilus. The
musk shrews classified in the speciose genus, Crocidura, are even more problematic. There is
also considerable uncertainty as to how many species of genets (Genetta) occur in south-central
Africa.
The precise taxonomic affinities of both Roan and Sable antelopes require unequivocal
elucidation, as does that of Blue wildebeest occurring in southwest and central Zambia. This
situation means that we cannot state with certainty the precise identities of those populations
found in SE Angola. These three antelope populations are currently perceived as no more than
representatives of populations described in South Africa in the 19th century, but this hypothesis
has yet to be tested by comparing all museum material together with molecular analyses. It is
impossible to name many populations of African antelopes precisely, let alone define the ranges
of described taxa with any confidence (Cotterill 2003a).
Higher level revisions of the Mammalia continue, and growth in interest in phylogenetic
systematics over the past two decades suggests that radical changes in the taxonomy of all
mammals will continue for some years to come, including the cats, Family Felidae (Wilson &
Reeder 1993) and the Bovidae.
6.2
SOURCES OF KNOWLEDGE
6.2.1
Historical Aspects
This section first summarises the main sources of knowledge on mammals (natural history
collections and publications), which constitute the main historical contributions in each of the
five countries. In many respects, the relevance of much of this knowledge, where credible,
increases with age, as shown by the information, albeit cursory and incomplete, on past
distributions of some species in the 19th century and the impacts of phenomena such as the
rinderpest pandemic of the 1890s.
The first published information on the mammals occurring in the region summarises
observations made by the first European explorers. Collections focused on large mammals, and
some publications included descriptions of new species (Spinage 1995). This literature also
records important historical events, especially the rinderpest pandemic. In the 1870s Selous
(1881, 1908) and Oates (1971) made the first collections of mammals from northern
Matabeleland. Gibbons (1898, 1904) crossed NW Botswana and SW Zambia both before and
after the rinderpest outbreak of 1896, and later wrote of the decimation of many populations of
antelope that had been exceedingly abundant on the Shesheke Flats and in the area that today lies
in the Chobe National Park. The importance today of this 19th century literature is that it is the
only written information available about species which have declined greatly (especially Puku)
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or become locally extinct (White rhinoceros) in the region. Particular cases are discussed in more
detail below.
6.2.2
Taxonomic Diversity and Surveys
Smithers (1971), Ansell (1978) and Smithers and Wilson (1979) are principal sources of
knowledge on which mammals occur in Botswana, Zambia and Zimbabwe, respectively. Hill
and Carter (1941) summarise the little that is known about the mammals of Angola, while the
mammals of the Caprivi Strip in eastern Namibia have been summarised by Griffin and Grobler
(1991) and Brown and Jones (1994). There has also been more recent collecting and surveys of
particular areas. This is discussed in the following country summaries.
Angola
The definitive review of the mammals fauna of Angola remains Hill and Carter (1941). They
included specimens and records of A.S. Vernay and H. Lang who collected across south-central
Africa. Specimens from the latter are preserved in the American Museum of Natural History,
New York (AMNH) and in the Field Museum of Natural History, Chicago (FMNH).
Figure 6.1 indicates the distribution of bat specimens across the whole country showing there is
virtually northing known from the eastern portion of Angola. This acute gap in knowledge also
applies to other mammal groups.
Figure 6.1. Distribution of collecting sites of all known bat specimens (Chiroptera) from Angola showing
an acute gap in knowledge, especially in the east of the country. Each marked quarter-degree square
represents at least one specimen. After Crawford-Cabral 1989.
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Botswana
The first major survey of mammals in the then Bechuanaland was that done by the Vernay-Lang
Expedition (1930-1931), which collected mammal specimens across the territory including parts
of Four Corners area. Collecting extended as far to the northeast as Kazungula in the then
Southern Rhodesia (Smithers 1971, F.P.D. Cotterill unpublished data).
The principal source of information resides in the large collection built up during the Botswana
Mammal Survey from 1964 to 1969, which involved the then Museums of Rhodesia in
collaboration with the Mammal Section of the Smithsonian Institution, Washington DC. The
latter provided fully-equipped teams over a period of three years (Smithers 1971). The
approximately 100,000 mammal specimens collected are preserved either in the Natural History
Museum, Bulawayo (NMZB) or in the National Museum of Natural History (USNM).
Multi-disciplinary expeditions by what is now the Natural History Museum of Zimbabwe began
in the 1960s as part of the Botswana Mammal Survey. These continued through to the mid-1980s
in collaboration with senior school natural history societies, when several trips focused on the
Okavango Delta. This material, much of which has yet to be formerly reported upon, represents
notable range extensions for some bat species (F.P.D. Cotterill, unpublished data). An important
collection of Chiroptera from the Okavango Delta by A. Archer (partially reported upon in
Archer 1977) is preserved in the Smithsonian in Washington DC, with just a few specimens in
the Royal Ontario Museum (ROM) in Toronto. Some comparatively rare species have
noteworthy ecological and life history data.
Namibia
Shortridge (1934) summarised the available information about the mammals of the then South
West Africa, including the Caprivi Strip. The Bernard Carp Expedition of 1947, centred on the
eastern section of the Caprivi Strip, involved biologists from the Kaffrarian Museum (King
Williamstown), the Transvaal Museum (Pretoria) and the then National Museums of Southern
Rhodesia. It was followed by an expedition by the Transvaal Museum (Haacke, Rautenbach &
Kemp 1971). Both expeditions resulted in some important mammal specimens (Meester 1973).
More recent biological studies in the Caprivi (notably Brown & Jones 1994) include an
assessment of the biogeography and conservation status of the mammals. There have been
ecological studies on a few particular species; collecting and surveys continue (M. Griffin, pers.
comm.).
Zambia
The definitive summary of knowledge remains the Mammals of Zambia (Ansell 1978), a reliable
and comprehensive source on the taxonomy and biogeography of all mammals occurring in
southwestern and central Zambia. Compared to elsewhere, the southwestern corner of the
country is poorly surveyed. Collections of small mammals in the Choma and Kalomo districts in
the 1990s provide important records of some bats.
Zimbabwe
A significant portion of the northwest corner of Zimbabwe is included within the Four Corners
area, including Hwange National Park the mammal fauna of which was surveyed by Wilson
(1975). This information and additional biogeographical records were collated by Smithers and
Wilson (1979). Since 1982, there have been no less than five multi-disciplinary expeditions
under the auspices of the Natural History Museum of Zimbabwe (Bulawayo) focusing on small
mammals along the Zambezi upstream of Victoria Falls, in the Gwayi valley, the Matetsi Safari
Area and in Kazuma Pan National Park.
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6.2.3
Ecological Studies
Angola
There are no known studies on the ecology of mammals occurring in SE Angola. This is a
glaring gap in our knowledge, underwritten by the nearly total lack of any biological specimens
from this area.
Botswana
Ecological studies of mammals in Botswana have focused on large ungulates and carnivores
(Hunter 1991). One exception is the monitoring of population dynamics of murids on the Chobe
floodplain (Sheppe & Haas 1981), a study that extended from June 1971 to August 1972 (Sheppe
1972). Simpson (1974a, 1974b, 1974c) carried out a detailed ecological and behavioural study
on bushbuck in the Chobe National Park, and also emphasized the adverse impacts of elephants
on habitats and biodiversity in riparian habitats along the Chobe River (Simpson 1978). The
ecological survey of NE Botswana by Child (1968) focused on large mammals including several
antelope species, notably Common waterbuck, Red lechwe, Puku (Child 1975, Child & Von
Richter 1969) and Tsessebe (Child, Robbel & Hepburn 1972). Games (1983) studied Sitatunga
in the Okavango in some detail. The social behaviour of Grey-footed baboons, Papio griseipes,
has been intensively studied in Moremi Game Reserve (Johnson 2003; Silk, Seyfarth & Cheney
1999). Some of these studies have formed part of control programmes of economically important
hosts of epizootic diseases, especially trypanosomiasis (Hunter 1991) and foot and mouth disease
(e.g. Falconer & Child 1975, Hunter 1991).
Elephants have been the subject of detailed study (e.g. Ben-Shahar 1996, Stokke 1999, Stokke &
Du Toit 2002), while studies on predators have included ones on Spotted hyaena in Chobe NP
(Cooper 1989, 1990), and on disease in large felids (Osofsky et al. 1996) and Lycaon pictus
(Alexander et al. 1996). The Botswana National Parks department has maintained long term
monitoring programmes censusing populations of large mammals within and around protected
areas. Recent data for antelopes were included in an assessment of the conservation status of the
antelopes of Botswana (Ross et al. 1998). There are currently several studies of predators and
large ungulates, including an exhaustive monitoring of elephant movements and ecology (M.
Chase, pers. comm.). A recent study by C.G. Hunter (1966) of regional changes of Buffalo
focused on the population common to NE Botswana and NW Zimbabwe, demonstrating seasonal
migrations in relation to predation, hunting pressure and resource availability.
Namibia
Important ecological and other studies on the mammals of the Caprivi are mostly in the form of
reports or ‘grey’ literature of limited distribution, such as that by Brown and Jones (1994).
Published studies include Child (1975), Williamson (1981), Griffin and Grobler (1991) and
Schlettwein et al. (1991). There have been regular annual censuses of large mammal populations
in protected areas (Mamili and Mudumu National Parks) in the Caprivi (Schlettwein et al. 1991).
Zambia
The major focus of ecological research in the Four Corners portion of Zambia has been on the
Kafue National Park. Studies looked especially at antelopes in the 1960s (e.g. Hanks 1969a;
Hanks, Stanley Price & Wrangham 1969), and on certain small mammals (Wrangham 1969).
There have been important studies on the ecology of large species (including population
dynamics), and these have been briefly reviewed in the management history of the Kafue NP
(Mwima 2001). Species studied in some detail include Crawshay’s waterbuck, elephant, Puku,
Blue wildebeest, Oribi and Red lechwe. Other ecological surveys in the Kafue assessed predation
(Mitchell, Shenton & Uys 1965) and animal movements, and the monitoring of large mammal
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populations using road and aerial counts (Mwima 2001). There has been little, if any, research on
mammals in SW Zambia since then, except for that by Tembo (1995).
Zimbabwe
The majority of ecological studies within the Four Corners area have been restricted to
populations of large mammals, mainly in Hwange National Park. Most were carried out in the
1960s and 1970s; earlier studies are cited by Wilson (1975). The species were selected because
of their relevance to ecology and management (elephant, hyaena) or conservation (White rhino,
Painted hunting dogs). Fergusson (1990), for example, reviewed the population dynamics of
Sable antelope, a key trophy species of management concern in Matetsi Safari Area.. Most
studies have focused on elephants (e.g. Williamson 1975a, 1975b) and were conducted under the
auspices of the Department of National Parks and Wildlife Management (DNPWM). They have
been allied with regular censuses of large mammals, notably in Matetsi (Fergusson 1990). An
important collation of research was presented at a conference at Hwange Safari Lodge in 1990;
unfortunately these proceedings were not published (F. Robertson, pers. comm.).
Behavioural and ecological studies of large carnivores in and around Hwange NP have received
more attention beginning in the 1990s. Subjects have included Black-backed and Side-striped
Jackals (Loveridge & MacDonald 2001, 2002), Lion (A. Loveridge, pers. comm.), Spotted
hyaena (Salnichi et al. 2001), and especially Painted hunting dogs (Davies 1993, Woodroffe &
Ginsberg 1999a,1999b, Rasmussen 1999, Girman et al. 2001). Hyaena and Painted hunting dogs
continue to be intensively studied, with a focus on the packs along the eastern and southern
boundaries and in the northwest (Sinamatella and Robins camps).
The ecology and population dynamics of elephants have been intensively studied since the
1960s, and censuses continue, which also monitor other large mammals. The research has been
allied with a revealing, long term taphonomic study of pachyderm mortality (Haynes 1993).
6.3
PRINCIPAL ASSEMBLAGES OF MAMMALS
The rich mammalian fauna of the Four Corners area owes its occurrence to several different
factors. The main one is the contribution from different faunal elements comprising species
associated with certain landscapes such as savanna woodlands (arid and mesic), grasslands,
rocky hills and gorges (rupicolous), and wetlands. This last category includes large rivers (such
as the Upper Zambezi), swamps (notably the Okavango) and riparian forests associated with
these permanent water bodies. Mammals characteristic of each of these principal faunal elements
are listed in Table 6.1.
The present-day habitat mosaic represents a transient stage when considered at evolutionary
scales; one has to consider the evolutionary history of the landscape. As discussed in more detail
below, climate change and drainage evolution through the Plio-Pleistocene have been major
determinants of the modern biodiversity of the area.
6.3.1
Kalahari Savanna
The distributional limits of many desert species (southwest arid endemics) corresponds to the
southern edge of the Okavango Delta and SW Makgadikgadi Pans. These include the gerbils,
Desmodillus auricularis and Gerbillurus paeba (the latter reaches the southern end of Hwange
NP), three small carnivores - Yellow mongoose, Cynictis penicillata, Black-footed cat, Felis
nigripes and Cape fox, Vulpes chama - and Springbok, Antidorcas melampus. Red hartebeest,
Alcelaphus cama rarely occurs to the east, with sporadic records from western Zimbabwe.
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Table 6.1. Mammal species that show an association with the principal faunal elements in the Four
Corners area.
Kalahari Arid
Savanna
Dry Savanna
Woodland
Otocyon megalotis
Bat-eared fox
Paracynictis selousi Epomops dobsoni
Selous mongoose
Dobson's fruitbat
Suricata suricatta
Suricate
Giraffa
camelopardalis
Giraffe
Mops niveiventer
Damaliscus lunatus
White-bellied free- Tsessebe
tailed bat
Lutra
maculicollis
Spotted-necked
otter
Cynictis
penicillata
Yellow mongoose
Tragelaphus
strepsicerus
Greater kudu
Papio kindae
Kinda baboon
Redunca arundinum
Reedbuck
Tragelaphus
spekei
Sitatunga
Felis nigripes
Black-footed cat
Damaliscus lunatus
Tsessebe
Genetta
angolensis
Angolan genet
Ourebia ourebi
Oribi
Kobus leche
Red lechwe
Hyaena brunnea
Brown hyaena
Aepyceros
melampus
Impala
Dendrohyrax
arboreus
Tree hyrax
Thryonomys
swinderianus
Greater canerat
Kobus vardoni
Puku
Alcelaphus cama
Red hartebeest
Raphicerus
campestris
Steenbok
Sigmoceros
lichtensteini
Lichenstein's
hartebeest
Pedetes capensis
Springhare
Dasymys
incomtus
Water rat
Antidorcas
marsupialis
Springbok
Raphicerus sharpei
Sharpe's grysbok
Heliosciurus
gambianus
Gambian sun
squirrel
Dendromus
melanotis
Grey climbing
mouse
Dendromus
mesomelas
Chesnut
climbing mouse
Xerus inaurus
Ground squirrel
Pronolagus
randensis
Jameson's red rock
rabbit
Anomalurus
derbianus
Scaly-tailed flying
squirrel
Dendromus nyikae
Climbing mouse
Dendromus
mystacalis
Lesser climbing
mouse
Zelotomys
hildegardeae
Hildegarde's rat
Arvicanthus
niloticus
Desmodillus
auricularis
Namaqua gerbil
Mesic Miombo
Woodland
Grassland
Wetland
Connachaetes
taurinus
Blue wildebeest
Pipistrellus
rueppelli
Rueppell's bat
Gerbillurus paeba
Hairy-footed
gerbil
Lemniscomys rosalia
Single-striped mouse
Zelatomys
woosnami
Woosman's desert
rat
Rhabdomys pumilio
Striped mouse
238
6.3.2
Mopane Savannas and Dry Miombo Woodlands
This faunal assemblage, which includes that in Baikiaea woodlands on Kalahari sands, accounts
for the majority of mammal species occurring in the area. Species with a southern savanna
distribution include Greater kudu, Tragelaphus strepsiceros, Sharpe’s grysbuck, Raphicerus
sharpei and Selous’ mongoose, Paracynictis selousi.
6.3.3
Mesic Miombo Woodlands
Mammals associated with the mesic miombo woodlands of central and northern Angola,
northern Zambia and Katanga in the Democratic Republic of Congo (DRC) only reach the
northern periphery of the study area in the Kafue NP. Examples include the Gambian sun
squirrel, Heliosciurus gambianus and Kinda baboon, Papio kindae. The Tree hyrax,
Dendrohyrax arboreus, also reaches its southern distribution in S Zambia (Ansell 1978).
The southernmost occurrences (Jeffery, Bell & Ansell 1989) of two forest antelopes (Yellowbacked duiker, Cephalophus silvicultor and Blue duiker, Philantomba monticola) reflect more
the availability of patches of evergreen thickets and gallery forest than mesic miombo woodland.
These two species illustrate (as does the Tree hyrax) the influence of evergreen dry forest and
thicket in augmenting species richness. The precise determinants for the southern limit of these
species within and among mesic miombo woodland are difficult to identify. It is likely that this
southern margin corresponds not only to these habitats but also to the former course of major
wetlands, especially that of the palaeo-Chambeshi and palaeo-Kafue rivers. The former extent of
the northeastern part of palaeo-Lake Makgadikgadi was probably an important determinant as
(judged by the alluvium of the Machili Flats) this lake stretched back to be contiguous with the
Kafue Flats (Cotterill 2003b).
6.3.4
Wetlands, Grasslands and Riparian Woodlands
Besides the dominance of wetlands, the overall habitat diversity in the Okavango Delta is of
great significance in supporting a rich mammal fauna. Riparian forests and termitaria are
particularly important microhabitats for small mammals. The southern portion of the study area
corresponds to a local hotspot in southern Africa, where over 40 species of grazing mammals
occur (Andrews & O’Brien 2000).
Grasslands in the area are either semi-arid habitats within Kalahari savanna, or are closely
associated with wetlands along edges of the principal rivers and swamps. Important examples are
Kazuma Pan and the Shumba Flats in NW Zimbabwe, utilized by grazing ungulates including
Blue wildebeest and Tsessebe. These attract large predators, notably Painted hunting dog and
Cheetah.
The richness of the small mammal fauna of these grasslands varies with respect to their
association with perennial water. Close proximity to perennial water increases species richness
(as in the Okavango and Linyanti). The shrews Crocidura occidentalis and Suncus lixus, are
associated with wetlands and moist grasslands, as are many rodents including Water rat Dasymys
incomtus, Groove toothed rat Pellomys fallax, vlei rats Otomys spp. and the climbing mice
Dendromus mystacalis and D. mesomelas.
6.3.5
Rocky (Rupicolous) Habitats
Rupicolous mammals constitute another significant component of the mammal fauna, although
rocky habitats are very patchy within the Four Corners area. The principal locations are in the
northern section of Hwange NP (especially the hills from Sinamatella northwards across the
Deka valley to the Gwayi and Zambezi). There are also rugged basalt hills and valleys
downstream of Victoria Falls, extending either side of the Batoka gorge. This young drainage
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system is characterised by numerous narrow gorges incised into the basalts and Karoo
sandstones. Patchy rupicolous habitat extends across NW Zambia along the northern edge of the
Gwembe rift. Granite outcrops are also found in parts of Choma District and in the Kafue NP.
The Tsodilo Hills west of the Okavango provide similar habitat.
This rocky habitat has a distinctive associated fauna, including two species of hyraxes, Procavia
capensis and Heterohyrax ruddi, Klipspringer Oreotragus oreotragus, Rock elephant shrew
Elephantulus myurus, Jameson’s red rock rabbit Pronolagus randensis, and several species of
rupicolous rodents. Notable examples are the rock dormouse Graphiurus platyops, Namaqua
rock rat Aethomys namaquensis, and Spiny mouse Acomys spinosissimus.
6.4
ECOLOGICAL AND EVOLUTIONARY PATTERNS AND DETERMINANTS
6.4.1
Ecological Determinants
In contrast to the eutrophic savannas of East Africa, much of the Four Corners area is semi-arid
savanna (miombo and Kalahari sand woodlands) on dystrophic soils. Hence the area supports a
comparatively low biomass of mammals (Coe, Cumming & Phillipson 1976, East 1984). As
judged from densities of raptor nests in Hwange NP, primary productivity has been interpreted to
be significantly greater on basalt soils as determined from the greater availability of small
vertebrates and mammals as prey (Hustler & Howells 1990). This difference from the Kalahari
sands could reflect other differences between the ecology of Kalahari sand and basalt habitats,
for example grasses growing on basalt soils are more palatable. It remains to be established if
primary production is in fact higher on the Kalahari sands (K. Dunham, pers. comm.).
The overall pattern of comparatively low mammal biomass changes in parts of the area,
especially in the E Caprivi and NE Botswana. The network of wetlands, especially distributaries
of the Okavango and Cubango rivers east of the Okavango Delta, are of great ecological
significance, especially where surface water persists for long periods.
6.4.2
Mammals as Ecological Determinants
The ecological engineering activities of Hippotamus are important in maintaining and expanding
aquatic habitats in the Okavango Delta, especially where they open up channels and thus
facilitate water flow between channels and permanent lakes. These activities extend the area of
permanent swamp (McCarthy, Ellery & Bloem 1998).
In addition to humans, elephants can be singled out as a major determinant of the functioning
and structure of savanna landscapes. This especially applies to their activities in modifying
habitat structure of woodlands (Cumming et al. 1997) and around localized water bodies
(Haynes 1993).
6.4.3
Landscape Evolution
Landscape evolution in the Four Corners area has been profound (see Moore, Chapter 2), in that
the drainage systems across south-central Africa have radically changed. This process began
before the Miocene, with especially significant alterations of the Upper Zambezi and Okavango
system through the Plio-Pleistocene. Of particular relevance was the existence of palaeo Lake
Makgadikgadi through the Pleistocene. Its huge area was maintained in part by run-off from the
present Upper Zambezi and the Okavango and Cubango rivers, which today flow into the
Okavango and Chobe. Their inflows were significant, but the largest endoreic tributary of palaeo
Lake Makgadikgadi was the palaeo-Chambeshi River, that flowed southwest from NE Zambia
parallel with the Muchinga Escarpment, to terminate in the Kalahari basin (Figure 6.2). The most
apparent signature of its importance is the vast alluvial deposits extending from NE Botswana
240
into SW Zambia that are contiguous with the present day Kafue Flats (Moore & Larkin 2001,
Cotterill 2003b).
Figure 6.2. Extent of alluvial deposits in south-central Africa. Data collated from geological survey maps
for Katanga, DRC and Zambia; extent of alluvium in NE Botswana modified from Moore & Larkin
(2001). Adapted from Cotterill (2003e) and Broadley & Cotterill (2003).
241
The physical evidence is complemented by geographical distributions of particular wetland
biota, for example vertebrates and Odonata, which occur in the drainage systems of both the
Chambeshi and Upper Zambezi-Okavango (see reviews in Timberlake 2000). Interesting
biogeographical patterns are evident in the otherwise anomalous discontinuities of many
vertebrates, including several mammals (Cotterill 2003a, 2003b; Broadley & Cotterill 2003). A
signature of this link is represented in distributions of those wetland species (including Sitatunga,
Puku and Crawshay’s waterbuck) that share a common southern distribution limit in the
Okavango or Upper Zambezi, and are confined to upstream of Victoria Falls. In this respect, the
distribution of the Spotted-necked otter Lutra maculicolis is also restricted to large wetlands. At
least in south-central Africa, this species appears to have an Upper Zambezi distribution as it is
absent from the Zambezi downstream of Katombora rapids. The last locality is noteworthy as
two species of Otter (the other being the Clawless otter Aonyx capensis) occur sympatrically
here.
The distribution pattern of lechwe is even more informative, as vicariance of the former palaeoChambeshi drainage in Zambia and SE Katanga is the most plausible explanation for speciation
in these aquatic antelopes. Within the area only Red lechwe is confined to the Upper Zambezi
drainage system, while the Kafue lechwe is endemic to the Kafue Flats where it occurred
marginally within the Kafue NP (discussed below). The present pattern points to a formerly more
contiguous distribution from the Okavango (palaeo-Makgadikgadi) extending northeast along the
palaeo-Chambeshi to SE Katanga and Bangweulu (Cotterill 2000b, 2003c). The geologically
recent connection of its wetlands with NE Zambia and Katanga is important in understanding the
affinities of the extant biodiversity of the Four Corners area.
As mentioned above, several mammals in mesic miombo savanna have similar southern margin
limits to their ranges, corresponding roughly to an axis extending southwest from the middle
Kafue River north of the Gwembe rift to Kazungula. This boundary corresponds to the former
course of the palaeo-Kafue and palaeo-Chambeshi rivers. Examples include Tree hyrax
Dendrohyrax arboreus, Scaly-tailed flying squirrel Anomalurus derbianus and Gambian sun
squirrel Heliosciurus gambianus. The relationships of some of these species, as shown in the
primates, awaits clarification (Cotterill 2003a, 2003b, unpublished data).
An interesting, but still only partly researched, example is the molerats, Cryptomys spp. A
hotspot of speciation is centred on the central Zambian plateau, extending west and southwest to
the Upper Zambezi, where at least two species of localized distribution occur within the Four
Corners area (Burda et al. 1999). The evolutionary history and taxonomy of these populations of
molerats in Zambia are the subject of current research (P. Van Daele, pers. comm.). The most
plausible mechanism for their vicariance involves geological determinants. It appears that
drainage history has interacted with climatic changes through the Plio-Pleistocene to drive
extensive speciation in these burrowing mammals. Likely mechanisms included changes in the
extent and patchiness of Kalahari sands and alluvium and the varying geographical barriers in
the form of large wetlands (Cotterill 2003b).
Tectonic activity which has been the ultimate causation of these geomorphological dynamics has
been along the southwest edge of the East African Rift system. Repeated episodes of crustal
flexure and vertical movements along NEE-SSW trending faults lines have broken and also
reconnected major rivers. Headwater capture (in particular, the Mid-Zambezi capture of the
Upper Zambezi) has been an especially significant mechanism. This was in turn an outcome of
uplift and faulting of the landscape by tectonic activity, and has occurred comparatively recently
in geological time. With respect to the evolution of the biota of the Four Corners area, radical
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alterations of the drainage system since the Miocene (i.e. over the past 5 million years) have
been profound (Moore & Larkin 2001; Moore, Chapter 2).
6.5
NOTABLE POPULATIONS OF LARGE MAMMALS, INCLUDING HISTORICAL
EXTINCTIONS AND RANGE CONTRACTIONS
This section draws attention to particular species of large mammals which occur in large
concentrations and/or are of conservation concern. Large populations, especially where they
occur in dense concentrations, are of great ecological and economic significance.
6.5.1
Carnivores
The predators of Chobe National Park are comparatively well-studied, and are well-known
internationally thanks to popular wildlife documentaries. This is especially true of lions and
spotted hyaenas.
Lion, Panthera leo
Lions are a keystone predator in the Four Corners area. They are a popular trophy species, with
significant quotas allocated each year in Matetsi and other hunting areas within northwest
Zimbabwe. The Hwange Lion Project (http://www.wildcru.org/index.htm), carried out under the
auspices of the Wildlife Conservation Research Unit, Department of Zoology, University of
Oxford, began in 1999 and has continued into 2003. The main researcher, A. Loveridge, has
been comparing the population and social dynamics of an exploited lion population with those of
the protected population in Hwange NP.
Painted Hunting Dog, Lycaon pictus
Childes (1988) and Davies (1993) provide the principal information available about Lycaon
pictus in NW Zimbabwe. Disease in this species has been investigated in Botswana (Alexander
et al. 1996). There is no current or recently published information on this species from other
countries within the area, but there have been two research and monitoring projects on the
species centred in Hwange NP. These have studied packs around Sinamatella and Main Camp,
and the latter population has continued to be monitored after Davies’ (1993) study. The Four
Corners area constitutes one of the most important protected areas in Africa for this highly
mobile and threatened predator.
6.5.2
Ungulates
Elephant, Loxodonta africana
The distribution of elephant has changed profoundly in south-central Africa since the early 19th
century, when it was widespread across the region (see Hoare, Chapter 14). Its modern
distribution reflects a long history of persecution by humans. Latterly, competition and conflict
between humans and elephants for habitat has increased, especially in the semi-arid areas of
Zimbabwe (Smithers 1983). Nevertheless, the Four Corners area supports one of the largest
populations in Africa, with high densities in NW Zimbabwe and in Chobe NP.
Black Rhinoceros, Diceros bicornis
Once widespread throughout the area, Black rhinos were nearly extinct in NE Botswana by the
late 1960s. Their range had been reduced greatly since the early 1800s, and the population was
much reduced by 1890. All recent records were concentrated in the northeast part of the country
(Smithers 1971), where they moved between Botswana and the eastern Caprivi, and they are
believed to have disappeared by 1974 (Skinner & Smithers 1990). The few remaining Black
rhinos in Caprivi (east of the Kwando River) were translocated to safer habitat in the early 1990s
(Brown & Jones 1994).
243
Historically, the black rhino was widespread in Zimbabwe, including in the northwest. Selous
(1908) recorded it in the Deka valley and also, alongside White rhino, in the Dete vlei. A very
few individuals persisted in the northern region of Hwange NP into the late 1950s. These were
complemented by translocations of individuals from the Middle Zambezi, beginning in 1962
(Herbert & Austen 1972). The Sinamatella region of Hwange NP was proclaimed an Intensive
Protection Areas (IPZ) in the late 1980s, and a significant number of Black rhinos were
translocated into this area from the Middle Zambezi valley. Declining support and effectiveness
of conservation in these IPZs is part of the recent crisis afflicting conservation of both species of
rhino in Zimbabwe (Du Toit 2002).
There are historical records that Black rhino occurred throughout SW Zambia within the present
study area (Ansell 1978), and the species was very common in parts of the southern region of
Kafue NP until it was poached to extinction in the 1980s. Black rhino presumably also occurred
in SE Angola. Previously, Ansell (1969) estimated that a total of 1200 Black rhino occurred in
all of Zambia at that time, with approximately 130 individuals in Kafue NP.
White Rhinoceros, Ceratotherium simum
Although extinct from the region many years ago, White rhino had previously been widespread,
certainly in W Zimbabwe (Selous 1908, Oates 1971, Herbert & Austen 1972) and across
Botswana and Namibia. Shortridge compiled a detailed history of its decline in south-central
Africa, and suggested it still occurred in southeast Angola, when they were "...'very rare' at
Lujana (S.E. Angola) in the Tschbombe bush" (Shortridge 1934: 425). White rhino had already
become extinct in Botswana between 1880 and 1890. The latter extirpation occurred within 20
years (Smithers 1971), and was part of the extermination of southern White rhino throughout its
range (except Zululand) in the first three decades of the 20th century (Rookmaaker 2002). The
species was reintroduced into the northeast of Hwange NP in 1966 and 1967, and the population
had increased to 32 individuals by 1972. There was some dispersal out of this protected area; at
least six individuals moved great distances west into Botswana and one was sighted near Khwaai
River in 1971 (Herbert & Austen 1972). Ironically, the population of White rhino in Hwange NP
was extirpated by poachers in 1993 (Rachlow 1998, Rachlow & Berger 1997). The northern
region (Sinamatella) of Hwange is one of four IPZs for rhino in Zimbabwe.
White rhino were also reintroduced into Chobe NP, where they have persisted. There is no
conclusive evidence that the species ever occurred in Zambia, although Ansell (1959) suggested
it may have occurred between the Upper Zambezi and Mashi rivers on the SE Angolan border,
including what is today the Sioma-Ngwezi National Park. If so, this species would also have
occurred in SE Angola.
Two pairs of rhino were introduced from South Africa in the early 1960s to Kafue and
Livingstone National Parks. The first attempt was unsuccessful, but a small population was
established in Livingstone (Ansell 1978), although poached to extinction by 1989. A second
population was established there in 1994 (K. Dunham, pers. comm). "It was quite unjustified to
attempt introduction of an alien species to the Kafue National Park, but it would not be
unreasonable to try and establish it eventually in the Sioma-Ngwezi National Park which lies
within its putative original range" (Ansell 1978: 50). It should be emphasized that no evidence
exists for former occurrence of White rhino east of the Zambezi in Zambia.
Burchell's Zebra, Equus burchelli
Burchell's zebra is widespread in the Four Corners area, and large populations have persisted in
NW Zimbabwe and NE Botswana. Local movements occur within the region, apparently in
response to patchy rainfall and the variable grass production that ensues (see Cumming, Chapter
244
13); Zebra are dependent on surface water. Large numbers of zebra and wildebeest moved
between NE Botswana (Makgadikgadi) and the Kalahari, but these migrations have been
disrupted by cattle fences (Smithers 1971).
This species is specifically distinct from the extinct Quagga, which only occurred south of the
Vaal River in South Africa (Grubb 1993). It appears that three subspecies of Burchell's zebra
occur in the study area; where their distributions intergrade in NE Botswana (Figure 6.3). These
are E. burchelli antiquorum from the west and south, integrading with E. b. chapmani in central
and NW Botswana and E. b. zambeziensis along the Zambezi River. The taxonomy of all zebras
is still uncertain, and awaits a thorough study of museum material (Grubb 1981, 1993). In its
inclusion of at least three genetically different populations of zebra, the Four Corners area
illustrates how a large protected area can conserve the genetic diversity of a complex of closely
related populations.
Figure 6.3. Distributions of the six subspecies of Burchell's zebra. The thick dotted line approximates the
southern limit to modern distribution of zebra in southern Africa, where Equus b. burchelli is extinct.
Modified from Grubb (1981).
Buffalo Syncerus caffer
This is an important species sold in trophy hunts, especially in Zambia and Zimbabwe. A recent
study of changes in the regional distribution of Buffalo across NE Botswana and NW Zimbabwe
demonstrated seasonal migrations in relation to predation, hunting pressure and resource
availability (Hunter 1996, 1999; Hunter & Kerley 1999). There is significant conflict with
humans in NE Botswana where buffalo exploit agricultural fields for dry season forage. The
Kazuma depression is an important movement corridor in this landscape, and vehicle strikes on
these animals are significant along the Kasane-Francistown road. Hunter & Kerley (1999)
245
suggested that this road should be modified to create an underpass (at least 80 m wide and 5 m
high) across the depression, which would reduce costs (including human deaths) and equally
maintain ecological functioning of the wetland across the larger landscape. Future threats to this
buffalo population include prospective agricultural development north of the Kazuma depression
in Botswana. Overall, the costs of such a scheme would be outweighed by the wildlife industry
in this region (Hunter & Kerley 1999). Buffalo also have a bearing on other migratory species,
including elephant and Tsessebe.
Roan Antelope, Hippotragus equinus
A relatively large population of Roan antelope has persisted in NE Botswana and was estimated
to number 1360 in 1997 (Ross et al. 1998). Kafue NP has been the stronghold for the species in
Zambia, "where it is locally quite common and probably numbers in the low thousands" (Jeffery
et al. 1989). Roan antelope have declined in all areas, notably Namibia (Schlettwein et al. 1991)
and Zambia, but the population in Botswana is comparatively healthy (Ross et al. 1998). The
species was formerly abundant around Kazuma Pan National Park in Zimbabwe and a breeding
population has been successfully established on Wildlive Game Farm, Kalomo in SW Zambia.
Sable antelope, Hippotragus niger
Healthy populations of Sable antelope have persisted in protected areas in Botswana, Zimbabwe
and Zambia. Like buffalo, this antelope ranks high in economic importance, both for game
ranching and as a trophy species. It has declined to very low numbers in Namibia (Schlettwein et
al. 1991, Brown & Jones 1994). Sable is an important trophy animal, and both trophy quality and
numbers declined in NW Zimbabwe through the 1980s (Fergusson 1990). Predation on calves
and juveniles appeared to be one factor, and another was the artificial selective pressure of
trophy hunting that targeted bulls with the largest horns (Ginsberg & Milner-Gulland 1994).
Waterbucks, Kobus ellipsiprymnus, K. crawshayi, K. penricei
At least three taxonomically distinct populations of waterbucks occur in the Four Corners area.
Common waterbuck, K. ellipsiprymnus, occurs across NE Botswana into NW Zimbabwe
(Smithers 1983), and Shortridge (1934) recorded this species in the East Caprivi. Crawshay’s
waterbuck, K. crawshayi, does not occur south or west of the Zambezi River (Ansell 1978,
Cotterill 2000b, 2003b).
Griffin and Grobler (1991) recorded the presence of a few "Defassa waterbuck" K. e. crawshayi
in the Zambezi area of the Caprivi between 1983-1987 (Cotterill 2000b). As originally stated by
Shortridge (1934), and following Ansell (1972, 1978), these are better allocated to K. penricei.
The species was considered extinct in the Caprivi by 1990, but the reason for its decline is not
known (Schlettwein et al. 1991). Its occurrence in Namibia was considered to represent vagrants
from Angola (Shortridge 1934, Skinner & Smithers 1990).
In many respects, this complex distribution of the three populations of waterbuck is congruent
with the three subspecies of zebras, with their introgressive zones traversing the area.
Puku, Kobus vardoni
This reduncine antelope was formerly widely distributed across the wetlands of Zambia,
including floodplains and dambos in the catchment of the Kafue and Upper Zambezi rivers, but
their numbers and range declined radically through the 20th century (Ansell 1978, Rodgers
1984). Puku are now nearly extinct in the Caprivi (Schlettwein et al. 1991) and are endangered
in Botswana. Stuart (1989) drew attention to their precarious status in southern Africa, with a
highly localized distribution on the Chobe floodplain - the Puku Flats. Overall, reduncine
populations have declined in the western Caprivi and certain species apparently occur marginally
246
(East 1989). Over the past 100 years, Puku numbers have declined greatly in the eastern Caprivi
(Smithers 1983, East 1989), and their status is marginal and precarious in Namibia (Schlettwein
et al. 1991). As of the 1980s, a relic population of approximately 50 animals persisted on the
Chobe floodplain in Botswana (Smithers 1983, East 1989). F.C. Selous first visited the Puku
Flats in 1874, and found them very numerous, but they were greatly disturbed by dense
settlement in 1876 by refugees fleeing political turmoil in Barotseland, from which they have
never recovered (Dollman 1921). Puku also declined to extinction on Impalila Island in the
Caprivi after its settlement in 1958 (Child 1968). Today, this relict population persists in
numbers of less than 100 (Ross et al. 1998). The status of Puku in Kafue NP appears to be stable
(Jeffery et al. 1989) although there was loss of habitat for puku and other aquatic antelopes with
the creation of Lake Itezhi-tezhi on the Kafue River when this impoundment was completed in
1975 (Mwima 2001).
Red Lechwe, Kobus leche and Kafue Lechwe, K. kafuensis
The Red lechwe, K. leche, was extremely numerous in the Four Corners region, especially in the
Okavango and Linyanti (Shortridge 1934, Smithers 1971). As of the late 1980s, a population of
over 4300 occurred in East Caprivi and 150-200 in West Caprivi (Linyanti Swamps and along
the Mashi and Upper Zambezi rivers). It has been estimated that several thousand red lechwe
occur in the Linyanti area of Botswana, and these migrate seasonally onto the Chobe floodplain
(Williamson 1981, East 1989).
The largest south-central African species, Kafue lechwe K. kafuensis, is endemic to the Kafue
Flats in central Zambia. It occurred throughout the Namwala Flats, from below the Itezhi-tezhi
gorge on the Upper Kafue to the Kafue Gorge on the mid-Kafue. Hydroelectric dams have since
been built in both gorges, with Itezhi-tezhi completed in 1975. Besides populations on private
game ranches, two Vulnerable populations of K. kafuensis are now restricted to the Blue Lagoon
and Lochinvar National Parks, divided by the Kafue Dam. As recorded by Pitman (1934), Kafue
lechwe occurred in the Four Corners area early in the 20th century, and small herds persisted
near Ngoma until at least December 1967 (Hanks 1969b). At that time, the species "only rarely
ranges across the national park boundary on the Nkala river, east of Ngoma" (Hanks 1969b).
Together with Black rhino, Kafue lechwe are one of the two species of mammals to have been
driven to extinction within the Kafue NP. These local extinctions of Kafue lechwe and Black
rhino in Zambia have occurred within the past three decades.
Sitatunga, Tragelaphus spekei
The Okavango and Linyanti Swamps support healthy populations of Sitatunga (Ross et al 1998),
with approximately 4000 in the Okavango alone. The species has been studied by Games (1983,
1984) and Williamson (1986). It appears to cope with high hunting pressure in much of its range,
but is threatened by drying out of its aquatic habitat caused by changes in hydrology (Ross 1992,
Ross et al. 1998). The population along the Upper Kafue on the eastern edge of the Kafue NP
was disrupted by the closure of the Itezhi-tezhi dam in the late 1970s, but a healthy population
occurs in the Busanga Swamps northwest of the Kafue NP and adjacent Game Management
Areas.
Tsessebe Damaliscus lunatus
Tsessebe antelope are endangered over much of their range, which has declined radically since
the beginning of the 20th century (Cotterill 2003c, 2003d). Northeast Botswana remains a
stronghold, where at least three major populations occur - in the Okavango, Savuti Marsh
(Chobe NP) and in eastern Chobe along the Zimbabwe border. These populations numbered over
9000 in 1997, although numbers have declined locally in the Linyanti area (Ross et al. 1998).
Originally widespread in the Caprivi, Tsessebe are now scarce in Namibia (Schlettwein et al.
247
1991, Brown & Jones 1994). They have declined greatly in W Zambia, and it is unclear whether
any still occur in the southwest in Sioma-Ngwezi NP and adjacent parts of Angola. The
population in NW Zimbabwe is migratory, moving in and out of Botswana and occurring
seasonally in Kazuma and Matetsi. A population of 100 animals on the Shumba plains in
Hwange NP had declined to less than 20 in 1997 (Anderson & Wilson 1998). The species is
considered endangered in Zimbabwe, where formerly healthy populations have declined
radically since 1999 following their widespread extirpation on commercial farms (Cotterill
2003d).
Selous (1881, 1908) observed that Tsessebe in the Caprivi calved in late September whilst the
herds on the Mababe Flats (today in Chobe NP) calved in November. The same difference in the
life histories of these populations was described by Child et al. (1972) in the late 1960s. This
subtle difference may reflect a former geographical barrier between these populations, the most
likely being palaeo-Lake Makgadikgadi in the late Pleistocene. However, the Linyanti and
Chobe wetlands could well maintain this isolation today (Cotterill 2003d).
Oribi, Ourebia ourebi
The range and populations of this small antelope have declined greatly across southern Africa. It
is very poorly represented in museum collections. Within the Four Corners area, Oribi occur in
Kafue NP and apparently still exist in SE Angola. An important population persists on game
ranches in the Choma District of Zambia (I. Bruce-Miller, pers. comm.). Their distribution is
always localized as they are dependent on open grasslands. The greater extent of this habitat is in
Zimbabwe but extends marginally into Botswana (East 1989; Ross et al. 1998).
Kazuma Pan supports an important population of Oribi (Anderson & Wilson 1998) that has
always been small (estimated at under 100 individuals in the early 1970s, Wilson 1975). This
population centred on the Kazuma depression should be considered Vulnerable. It would appear
to represent O. o. rutila Blaine 1922, and is thus distinct from the species elsewhere in
Zimbabwe, which was referred by Ansell (1972) to O. o. hastata (Peters 1852). The taxonomy of
Oribi requires thorough review with respect to the many scattered populations; their
phylogeography is currently being studied (J. Jansen Van Vuuren, pers. comm.).
6.6
WATER AVAILABILITY AND MOVEMENTS
The semi-arid climate of the Four Corners area makes for stochastic ecological conditions. In
particular, surface water is a very patchy and ephemeral resource, its availability being
determined by rivers and distributaries and episodes of rainfall which fill pans and pools. The
drainage system in the Matetsi Safari Area and some northern areas of Hwange on basalt is more
perennial. Surface water persists in the pools of the larger rivers (Bumbusie, Deka and Matetsi),
which dry out only during severe droughts. There are also persistent springs and seeps in these
landscapes. The long dambos (notably the Dete Vlei) in the catchments of the Gwayi River on
Kalahari sand are particularly important features of the landscape along the eastern boundary of
Hwange NP. Their heavy use by large mammals was first recorded by Selous (1881, 1908).
Much of SW Zambia, much of NE Botswana and SE Angola lies on Kalahari sand; surface water
is an especially and patchy resource. As has been shown in Hwange NP, the artificial supply of
surface water creates a complicated situation of costs and benefits.
It appears that the movement of large mammals was formerly an important phenomenon between
NE Botswana and the Central Kalahari (Child 1972, Smithers 1971, Cumming in Chapter 13).
This occurred in response to the availability of key resources, especially fresh grass and water,
the availability of which is determined primarily by variation in rainfall. The occurrence of
248
certain species across vast regions of the area can be described as transient as these large
mammals move in relation to surface water availability. Although many mammals possess
adaptations to prolonged water stress, the crux is in drier years when larger mammals move to
available water or die (Lovegrove 1993). Management of surface water supplies is a critical issue
in biodiversity conservation (Child 1972). It governs the distribution of elephants on a seasonal
and daily cycle, especially as small calves struggle to commute long distances between feeding
areas and drinking sites. As confirmed by Haynes (1993), the geography of water points
becomes the major determinant of elephant density and habitat use when water is seasonally
limiting. The resultant impacts on vegetation and terrestrial biodiversity are marked (Cumming
et al. 1997).
6.7
CONSERVATION STATUS AND CONCERNS
6.7.1
Threatened Species and Habitats
Ten (or 5%) of the mammals occurring in the Four Corners area are classified as of global
conservation concern (Appendix 6.1). Reasons for the vulnerability and declines of these species
are complex. Three can be singled out: (i) diseases, (ii) direct depredations by humans, and (iii)
loss of habitat. The impacts of unsustainable human offtakes of larger species coupled with
landscape changes are illustrated in the Caprivi where many wetland species are threatened or
are already locally extinct (Griffin & Grobler 1991).
The categories of conservation status are the officially recognized designations, which are
misleading when one comes to consider many species occurring in the area. Species such as
Oribi are vulnerable in all five countries, and except for parts of the Kafue NP, occur in only
small and isolated populations. It and many other species are either Rare and/or Vulnerable.
Furthermore, some species have declined significantly in some parts of their range (e.g. Tsessebe
in Caprivi and Zimbabwe) although they may exist in healthy populations elsewhere within the
area (Botswana in the case of Tsessebe). Many species of small mammal appear at risk of habitat
changes driven by damage from elephants and humans, shown in N Zimbabwe for tree-roosting
bats (Cumming et al. 1997).
The existing network of protected areas covers a wide range of the assemblages listed above;
including wetlands, savannas and rupicolous habitats. It is not possible in this review to carry out
a thorough gap analysis of how adequately protected mammal faunas are in terms of
distributions of species with respect to habitats. A significant gap in its own right is the patchy
knowledge of biodiversity. Patches of evergreen thickets and dryland forest are especially underrepresented in the existing network of protected areas. Hence the Tree hyrax Dendrohyrax
arboreus cannot be expected to persist over the long term within the area.
6.7.2
Diseases
The most infamous pandemic that decimated many bovids was the rinderpest which was
widespread across Africa and greatly reduced many species in the region in 1896 (Gibbons 1898,
1904, Spinage 1962). Carnivore populations are susceptible to particular diseases, notably viral
infections (Osofsky et al. 1996). Epidemics have greatly reduced populations of Lion and
Painted hunting dogs (Woodroffe, Ginsberg & Macdonald 1997; Creel & Creel 2002) in East
Africa. Feral and domestic dogs are a significant factor in the epidemology of rabies in parts of
Africa, and the increased contact between wild carnivores and dogs has extirpated populations of
the former (Alexander et al. 1993, 1996).
Certain murid rodents are carriers of the plague bacillus, Yersinia pestis, among many pathogens
harmful to humans. The principal vectors are the House rat Rattus rattus, Multimammate mouse
249
Mastomys natalensis and, to a lesser extent, Bushveld gerbil Tatera leucogaster. The
Multimammate mouse is a commensal species (De Graaf 1981, Smithers 1983). A major
outbreak of bubonic plague occurred in Ngamiland in 1944, apparently associated with high
flood levels in the Okavango delta (Davis 1946), and the disease is now endemic in the region
with sporadic outbreaks (Smithers 1983). In addition to the more apparent diseases such as
bilhzaria and malaria, bubonic plague needs to be considered in the development of urban
centres and tourist resorts.
6.7.3
Introductions of Threatened And Extinct Species
Translocations of the two species of rhinoceros were described earlier. If protected areas in the
Caprivi and elsewhere in the Four Corners area can be secured, than large mammals can be
introduced. Candidates include Penric's waterbuck into the Caprivi, and Black and White rhinos
into Sioma-Ngwezi NP in Zambia.
Any future translocations of organisms have to consider carefully the taxonomic and genetic
affinities of the populations being introduced. This especially applies to ungulates. Introductions
of mammals of populations from elsewhere in Africa (seen in the translocation of West African
Roan antelope into South Africa, Matthee & Robinson 1999) is tantamount to introducing an
alien species into an ecosystem. It is undesirable that short-term commercial concerns should
override scientific and ecological realities (Castley, Boshoff & Kerley 2001). This concern is
magnified in the Four Corners area for two reasons. One is the peripatric distributions of closely
related species and populations, as shown by Burchell's zebra and waterbucks, a situation that
dictates the need for thorough appraisals of which organisms are translocated where. Secondly,
subtle differences between apparently very similar populations have evolved (for example, the
differences in life histories of the Tsessebe in NE Botswana compared to those in the Caprivi),
further emphasising the need to recognize that such populations have evolved adaptations to their
particular environments.
6.8
MONITORING AND FUTURE STUDIES
Monitoring of mammal populations and assemblages is a complex subject. Many of these issues
and techniques are detailed and discussed thoroughly by Wilson et al. (1996) and Jachmann
(2001). Large mammal populations have been monitored within the principal protected areas of
the Four Corners area for several decades. Considered overall, these efforts have been
fragmentary and employed many different techniques and methods within the main themes of
aerial, vehicle and walked transects. Surveys have also used either total counts of a selected
population or indices of abundance. Censuses of elephants in Botswana and Zimbabwe are
considered relatively reliable, and this is the most feasible (and often the only) species which can
be counted from aircraft, especially in wooded savanna (Jachmann 2001).
6.8.1
Monitoring Large Mammals of High Conservation Concern
Overall estimates of small populations are invariably not sufficient as individuals need to be
distinguished. Within the study area, costs and human resources decree that the most feasible
approach is for game scouts to monitor animal activity (including usage of home ranges and key
resources). Monitoring of large carnivores has relied on an observer's intimate knowledge of
individuals, using photographs of pelage patterns (e.g. Painted hunting dog, G. Rasmussen, pers.
comm.). A similar approach was utilized for Leopard in East Africa (Miththapala et al. 1989).
Specialized techniques are available to monitor mammals of high conservation concern such as
rhinos using genetic signatures of individuals (microsatellite markers). This is feasible only for
relatively small, highly dispersed populations (Cunningham et al. 2001). Techniques based on
250
automated analyses of footprints of black rhinos (as in Sinametella IPZ in Hwange by Jewell,
Alibhai & Law 2001) are of dubious application, unless applied by experts in bushcraft (game
scouts), who in any case routinely distinguish between individuals by eye at a glance.
6.8.2
Ecological Studies
The mammal fauna might appear well known, but all we know about the majority of species is
that they occur in the area, and nothing more. This deficiency is especially true of nearly all the
small mammals. One feasible approach to remedy this gap is to carry out intensive markrecapture studies of small mammal assemblages in representative habitats. Such studies should
be for at least two years, and employ standardized, comparable methods. The localities studied
by Sheppe and Haas (1981) in Chobe should be revisited.
The primary objective of all these studies should be to establish the composition (species
diversity) of the mammal assemblages, while species of economic, ecological and conservation
concern can be singled out for more focused study. An important lesson from past research is to
frame these studies within the context of the overall landscape, in which biodiversity is
comphrehensively characterised (Cotterill 1995).
6.8.3
Biodiversity Surveys, Monitoring and Taxonomic Research
Southeast Angola is the obvious priority for a thorough survey of its biodiversity, which should
include all vertebrates and not just mammals. Nevertheless, major gaps remain in all countries,
as seen in the Caprivi where the presence of many listed species (Brown & Jones 1994) is based
on extrapolation from neighbouring surveyed habitats and suspected knowledge of the habitat
associations of a particular species. Only properly organized biodiversity surveys, which collate
comprehensive biological collections, will remedy these deficiencies.
A more detailed research plan for the Four Corners area should consider the inauguration of a
long-term and on-going biodiversity survey. Such a programme needs to choose target taxa
(small mammals are an obvious candidate) under the direction of professional biologists with
survey experience. A model is the Botswana Mammal Survey, in which conservation agencies
and numerous individuals made substantial contributions. The aim should be to build
comprehensive biological collections whose specimens are accompanied by detailed ecological
data. Conservation authorities need to be shown the importance of such a long term, multidisciplinary survey. One possible approach is to insist that safari operators with leases on state
land participate in biodiversity surveys. If this became a mandatory requirement, it would do
much to improve the knowledge of hunters and guides if they collected specimens of organisms
besides charismatic large mammals.
A major obstacle to perform biodiversity surveys efficiently is the so-called Taxonomic
Impediment (Hoagland 1996, Mims 2003). This is a global problem and afflicts virtually all
museums and herbaria. There are too few taxonomic experts to identify specimens, and these
posts are not being supported. The integrity of many of the world's natural history collections are
in severe jeopardy. In this respect, one also has to acknowledge that the responsible agencies in
the region, especially herbaria, National Museums of Botwana and Namibia, the Livingstone
Museum and the Natural History Museum of Zimbabwe, will require major improvements in
professional capacity to preserve and classify the large collections that will result from a full
biodiversity survey. The key deficiency is insufficient taxonomic expertise, therefore funding for
such a survey will have to include the cost of capacity building. All nations participating in the
Four Corners TFCA initiative have signed and ratified the Convention on Biological Diversity
which requires its members to survey their biodiversity scientifically.
251
The collation of comprehensive biological collections requires digital catalogues of their
specimens and attributes as a prerequisite to their management, and especially to support
research based on their attributes. These then provide valuable knowledge to multifarious queries
on environmental concerns. A suite of manuals and electronic databases are particularly suited to
improve the efficiency with which the primary data produced by biodiversity surveys can be
synthesized (see Cotterill 1995). There are, however, no short shortcuts around the extinction of
taxonomic resources (Cotterill 2002, Mims 2003).
6.9
MAMMAL CONSERVATION VALUES OF THE FOUR CORNERS TFCA
The occurrence of around 200 species of mammals in the Four Corners area shows the influence
of habitat hetereogeneity, which in turn is a function of the complex evolutionary history of the
landscapes of south-central Africa. The underlying reason for this diversity is the extremely large
size of the landscape enclosed within this international conservation area. With respect to
conservation of mammals, the Four Corners TFCA provides several benefits, which also confer
significant costs for management.
Being a large area, the TFCA conserves genetic heterozygosity, as seen with many vertebrates
where complexes of recently diverged populations or closely related species occur within the
region. The inclusion of hybrid zones (as for zebras in NE Botswana, and also for Kinda and
Grey-footed baboons in Zambia) is of equal value. These regions of active evolutionary change
could provide potential buffers against habitat changes and other disturbances, especially
diseases. Climatic change is an especially important consideration. One can argue that
conservation of a species at the edge of its range (where they may be declining or are
Vulnerable), is not worth the expense. Examples within the area include the two forest duikers.
These highly threatened species may not persist without a large investment in anti-poaching, and
perhaps will require maintenance and even restoration of their habitat. Can such an investment
be justified? There are at least three sound reasons why it is important to conserve these less
conspicuous and/or marginal habitats and species:
(a)Peripatric Diversity and Intraspecific Variation. Considered across its entire range, a
species at the edge of its range may possess adaptations and genetic diversity which are not
present in source populations or in the central population of the species. Based on the theory of
population genetics (Meffe et al. 1997), genetic novelties may also accumulate more rapidly and
become fixed in such smaller, peripheral populations where organisms are likely to encounter
novel environmental challenges.
(b)Habitat Hetereogeneity. Habitats which constitute islands in the Four Corners TFCA are
indeed geographically insignificant, marginal or very isolated, but they raise the beta and gamma
diversity across the landscape. In addition to small hills and similar rupicolous habitats, one
pertinent example is the evergreen thickets and forests in SW Zambia, including riparian fringes
along the Kafue and Lufupa rivers in the Kafue NP. Another example is the islands of rocky
habitat scattered across SW Zambia and in the Caprivi (e.g. Mushangara area).
(c)Future Climatic and Habitat Change. The edge effects associated with peripheral
populations, in which local adaptations may have evolved in demes, may predispose such species
and patchy habitats to accommodate climate change.
All these issues, both in this section and the previous discussion of monitoring and research,
need to be evaluated and accommodated in a comprehensive research and management plan for
the Four Corners TFCA. Any such plan will obviously have to prioritize investments in research
252
and management with respect to conservation problems, threats to biodiversity and ecological
integrity and, especially, gaps in knowledge.
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260
Appendix 6.1. Mammals occurring in the Four Corners area, by country.
Data from Ansell (1978), East (1989), Hill & Carter (1942), Smithers (1971), Smithers & Wilson (1979).
Taxonomy follows Wilson & Reeder (1993), with alterations from Groves (2001) and Kearney et al.
(2002) for primates and vesper bats, respectively. Records for Namibia are conservative, being restricted
to material evidence only; Brown & Jones (1994) list many species which are expected to occur, indicated
by '?'. Conservation status and categories follow IUCN Red Data list 2003 (http://www.redlist.org),
categories CR = Critical, EN = Endangered, VU = Vulnerable, DD = Data Deficient, LR/cd = Low Risk,
conservation dependant, LR/nt = Low Risk, not threatened. These have been amended for the Four
Corners area, where R = Rare, V = Vulnerable, Ex = Extinct, and apply to populations which occur
marginally in the region or are confined to patchy habitat.
Taxon
Conserv.
status
Ang Bot
Nam Zam Zim
Order INSECTIVORA (shrews and hedgehogs)
Family Soricidae
Suncus lixus (Thomas 1898) Greater dwarf shrew
LR/nt R
+
?
+
Crocidura cyanea (Duvernoy 1838) Reddish-grey musk shrew
DD
+
+
+
Crocidura flavescens (Geoffory 1827) Greater musk shrew
VU B1+2c
Crocidura fuscomurina (Heughlin 1865) Tiny musk shrew
LR/nt
Crocidura silacea Thomas 1895 Lesser grey-brown musk shrew
DD
Crocidura hirta Peters 1852 Lesser red musk shrew
LR/nt
Crocidura mariquensis (A.Smith 1844) Swamp musk shrew
+
+
+
+
+
+
+
+
+
+
+
+
+
DD
+
+
+
+
Crocidura occidentalis (Pucheran 1855) Giant musk shrew
LR/nt
+
+
+
+
Crocidura turba Dollman 1910 Dollman's musk shrew
DD
+
Family Erinaceidae
Atelerix frontalis (A.Smith 1831) Southern African hedgehog
LR/nt V
+
+
Order CHIROPTERA (bats)
Family Pteropididae
Epomophorus crypturus Peters 1852 Peters' epaulatted fruit bat
LR/nt
+
+
+
+
+
Epomophorus gambianus Gray 1870 Gambian epaulatted fruit bat
LR/nt
+
+
+
+
+
Epomophorus wahlbergi (Sundevall 1846) Wahlberg's epaulatted
fruit bat
LR/nt
+
+
?
+
+
Epomops dobsoni (Bocage 1889) Dobson's fruit bat
LR/nt R
+
?
+
+
Eidolon helvum (Kerr 1792) Straw-coloured fruit bat
LR/nt
+
+
+
+
Rousettus aegyptiacus (E.Geoffroy 1810) Egyptian fruit bat
LR/nt
+
+
+
+
+
+
+
+
+
Family Emballonuridae (tomb bats)
Taphozous mauritianus E.Geoffroy 1818 Mauritian tomb bat
LR/nt
+
+
Taphozous perforatus E.Geoffroy 1818 Egyptian tomb bat
DD
+
+
Tadarida aegyptiaca (E.Geoffroy 1818) Egyptian free-tailed bat
LR/nt
+
+
Tadarida fulminans (Thomas 1903) Madagascar large free-tailed
bat
LR/nt
Tadarida ventralis (Heughlin 1861) Giant African free-tailed bat
DD
Mops midas (Sundevall 1843) Midas free-tailed bat
LR/nt
+
?
+
Mops condylura (A.Smith 1833) Angola free-tailed bat
LR/nt
+
+
+
Family Molossidae (free-tailed bats)
+
+
261
Taxon
Conserv.
status
Ang
Bot
Nam Zam Zim
Mops niveiventer Cabrera & Ruxton 1926 White-bellied free-tailed
LR/nt
bat
+
Chaerephon ansorgei (Thomas 1913) Ansorge's free-tailed bat
LR/nt
?
+
+
Chaerephon bivittata (Heughlin 1861) Spotted free-tailed bat
LR/nt
+
+
Chaerephon chapini (J.A.Allen 1913) Long-crested free-tailed bat
DD
+
+
Chaerephon pumila (Cretzschmar 1830) Little free-tailed bat
LR/nt
+
+
+
+
Chaerephon nigeriae Thomas 1913 Nigerian free-tailed bat
LR/nt
+
+
+
+
Family Hipposideridae (leaf-nosed bats)
LR/nt
Cloeotis percivali Thomas 1901 Trident leaf-nosed bat
DD
Hipposideros caffer (Sundevall 1846) Sundevall's leaf-nosed bat
LR/nt
+
+
+
+
+
Hipposideros commersoni (E.Geoffroy 1813) Commerson's leafnosed bat
LR/nt V
+
+
+
+
+
+
Family Rhinolophidae (horseshoe bats)
Rhinolophus clivosus Cretzschmar 1828 Geoffroy's horseshoe bat
LR/nt
?
+
Rhinolophus darlingi Andersen 1905 Darling's horseshoe bat
LR/nt
+
+
+
+
Rhinolophus fumigatus Rüppell 1842 Rüppell's horseshoe bat
LR/nt
+
+
+
+
Rhinolophus hildebrandti Peters 1878 Hildebrandt's horseshoe bat
LR/nt
?
+
+
Rhinolophus landeri Martin 1838 Lander's horseshoe bat
LR/nt
+
+
Rhinolophus denti Thomas 1904 Dent's horseshoe bat
LR/nt
+
Rhinolophus simulator Andersen 1905 Bushveld horseshoe bat
LR/nt
?
Rhinolophus swinnyi Gough 1908 Swinny's horseshoe bat
LR/nt R
+
+
Family Nycteridae (slit-faced bats)
Nycteris hispida (Schreber 1774) Hairy slit-faced bat
DD
Nycteris macrotis Dobson 1876 Large-eared slit-faced bat
LR/nt R
Nycteris thebaica E.Geoffroy 1813 Egyptian slit-faced bat
LR/nt
Nycteris woodi K.Andersen 1914 Wood's slit-faced bat
LR/nt R
+
+
+
?
+
+
?
+
+
+
+
+
+
Family Vespertilonidae (vesper bats)
Miniopterus schreibersi (Kuhl 1819) Schreiber's clinging bat
LR/nt
+
?
+
+
Laephotis botswanae Setzer 1971 Botswana long-eared bat
LR/nt R
+
+
+
+
Neoromicia rendalli (Thomas 1889) Rendall's serotine bat (ex.
Eptesicus)
DD
+
?
+
Neoromicia capensis (A.Smith 1929) Cape pipistrelle (ex
Eptesicus)
LR/nt
+
+
+
Neoromicia melckorum (Roberts 1919) Melck's pipistrelle (ex
Eptesicus)
DD
+
?
Neoromicia somalicus (Thomas 1901) Somali pipistrelle (ex
Eptesicus)
LR/nt
+
+
+
+
Hysugo anchietai (Seabra 1900) Anchieta's pipistrelle (ex
Pipistrellus)
VU A2c
+
+
?
+
+
Pipistrellus rusticus (Tomes 1861) Rusty bat
LR/nt
+
+
+
+
+
Pipistrellus kuhlii (Kuhl 1819) Kuhl's pipistrelle
LR/nt
+
?
+
+
Pipistrellus rueppelli (Fischer 1829) Rueppell's bat
LR/nt
+
+
+
+
+
+
+
262
Taxon
Conserv.
status
Neoromicia nanus (Peters 1852) Banana bat
LR/nt
Pipistrellus sp.
DD
Glauconycteris variegata (Tomes 1861) Butterfly bat
LR/nt
Scotophilus viridis (Peters 1852) Lesser house bat
LR/nt
Scotophilus dinganii (A.Smith 1833) Yellow house bat
LR/nt
Scotophilus sp. House bat
DD
Nycteicinops schlieffeni (Thomas & Wroughton 1908) Schlieffen's
bat
LR/nt
Ang Bot
+
Nam Zam Zim
?
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Kerivoula argentata Tomes 1861 Damara woolly bat
DD
+
?
Kerivoula lanosa (A.Smith 1847) Lesser woolly bat
DD
+
?
+
+
Order PRIMATA (primates)
Family Lemuridae
Galagoides moholi A.Smith 1836 Lesser nightape
LR/nt
+
Otolemur crassicaudatus (E.Geoffroy 1812) Thick-tailed bushbaby LR/nt
+
Family Cercopithecidae
Cercopethicus aethiops (Linnaeus 1758) Vervet monkey
LR/nt
+
+
+
+
+
Papio griseipes Pocock 1911 Gray-foot (chacma) baboon
LR/nt
+
+
+
+
+
Papio kindae Lönnberg 1919 Kinda (Katanga) baboon
LR/nt
+
Order CARNIVORA
Family Viverridae
Civettictis civetta (Schreber 1776) African civet
LR/nt
+
Genetta angolensis Bocage 1882 Angolan genet
DD
+
Genetta genetta Linnaeus 1758 Small-spotted genet
DD
+
Genetta maculata (Gray 1830) Large-spotted genet (ex. G. tigrina)
+
+
+
+
+
+
+
+
+
+
DD
+
+
+
+
Aonyx capensis (Schinz 1821) Clawless Otter
LR/nt
+
+
+
+
Lutra maculicollis Lichtenstein 1835 Spotted-necked otter
VU A1c
+
+
+
M
Mellivora capensis (Schreber 1776) Honey badger
LR/nt
+
+
+
+
Poecilogale albinucha (Gray 1864) African striped weasel
LR/nt R
*
+
+
M
Ictonyx striatus Perry 1810 African striped polecat
LR/nt
+
+
+
+
+
+
Family Mustelidae
+
+
Family Herpestidae
Suricata suricatta (Schreber 1776) Suricate
LR/nt
+
Paracynictis selousi (De Winton 1896) Selous' mongoose
LR/nt
+
+
Cynictis penicillata (G.Cuvier 1829) Yellow mongoose
LR/nt
+
?
Herpestes ichneumon (Linnaeus 1758) Large grey mongoose
LR/nt
+
+
+
+
+
Galerella sanguineus (Rüppell 1836) Slender mongoose
LR/nt
+
+
+
+
+
Rhynchogale melleri (Gray 1865) Meller's mongoose
LR/nt
Ichneumia albicauda (G.Cuvier 1829) White-tailed mongoose
LR/nt
+
+
+
+
+
Atilax paludinosus (G.Cuvier 1829) Water mongoose
LR/nt
+
+
+
+
+
Mungos mungo (Gmelin 1788) Banded mongoose
LR/nt
+
+
+
+
+
+
+
+
263
Taxon
Conserv.
status
Ang
Bot
Nam Zam Zim
Helogale parvula (Sundevall 1846) Dwarf mongoose
LR/nt
+
+
+
+
+
Family Canidae
Otocyon megalotis (Desmarest 1822) Bat-eared fox
LR/nt
+
+
Canis adustus (Sundevall 1846) Side-striped jackal
LR/nt
+
+
Canis mesomelas (Schreber 1778) Black-backed jackal
LR/nt
+
+
Lycaon pictus (Temminck 1820) Painted hunted dog
EN C1
+
+
+
+
Vulpes chama (A.Smith 1833) Cape fox
LR/nt
+
+
+
+
+
+
Family Felidae
Caracal caracal (Schreber 1776) Caracal
LR/nt
+
+
+
+
+
Felis lybica Forster 1780 African wild cat
LR/nt
+
+
+
+
+
Felis nigripes Burchell 1824 Black footed cat
VU C2a(i)
Leptailurus serval (Schreber 1776) Serval cat
LR/nt
+
+
+
+
+
Panthera leo (Linnaeus 1758) Lion
VU C2a(i)
+
+
+
+
+
Panthera pardus (Linnaeus 1758) Leopard
LR/nt V
+
+
+
+
+
Acinonyx jubatus (Schreber 1775) Cheetah
VU C2a(i)
+
+
+
+
+
Crocuta crocuta (Erxleben 1777) Spotted hyaena
LR/nt
+
+
+
+
+
Hyaena brunnea Thunberg 1820 Brown hyaena
LR/nt
+
+
+
Proteles cristatus (Sparrmann 1783) Aardwolf
LR/nt
+
+
+
+
+
LR/nt
+
+
+
+
+
EN A1b
+
+
+
+
+
+
Family Hyaenidae
+
Order TUBULUDENTATA
Orycteropus afer (Pallas 1776) Antbear, Aardvark
Order PROBOSCIDEA
Loxodonta africana (Blumenbach 1797) Savanna elephant
Order HYRACOIDEA
Dendrohyrax arboreus (A.Smith 1827) Tree hyrax
LR/nt V
+
Heterohyrax ruddi (Wroughton 1910) Yellow-spotted hyrax
LR/nt
+
Procavia capensis (Pallas 1766) Rock hyrax
LR/nt
+
Order PERISSODACTYLA
Diceros bicornis (Linnaeus 1758) Black rhinoceros
CR A2abc
E
E*
E
E
R
Ceratotherium simum (Burchell 1817) White rhinoceros
NT
E
E
E
R
R
Equus burchelli (Gray 1824) Burchell's zebra
DD
+
+
+
+
+
Potamochoerus porcus (Linnaeus 1758) Bushpig
LR/nt
+
+
+
+
+
Phacochoerus aethiopicus (Pallas 1776) Warthog
LR/nt
+
+
+
+
+
Hippopotamus amphibius Linnaeus 1758 Hippopotamus
LR/cd V
+
+
+
+
+
LR/cd
+
+
+
+
+
Family Suidae
Order ARTIODACTYLA
Family Giraffidae
Giraffa camelopardalis angolensis Lydekker 1903 Angolan giraffe
Family Bovidae
Tribe Tragelaphini
264
Taxon
Conserv.
status
Ang Bot
Nam Zam Zim
Syncerus caffer (Sparrman 1779) Buffalo
LR/nt
+
+
+
+
+
Tragelaphus strepsicerus (Pallas 1776) Greater kudu
LR/nt
+
+
+
+
+
Tragelaphus scriptus (Pallas 1776) Chobe bushbuck
LR/nt
+
+
+
+
+
Tragelaphus spekei Speke 1863 Sitatunga
LR/nt V
+
+
+
+
+
Taurotragus oryx (Pallas 1776) Eland
LR/nt
+
+
+
+
+
Tribe Cephalophini
Cephalophus silvicultor (Afzelius 1815) Yellow-backed duiker
LR/cd R
+
Philantomba monticola (Thunberg 1789) Blue duiker
LR/cd R
+
Sylvicapra grimmia (Linnaeus 1758) Grey duiker
LR/nt
+
+
+
+
+
Redunca arundinum (Boddaert 1785) Common reedbuck
LR/cd
+
+
+
+
+
Kobus crawshayi P.L.Sclater 1894 Crawshay's waterbuck
LR/cd
Kobus ellipsiprymnus (Ogilby 1833) Common waterbuck
LR/cd
+
+
+
+
Kobus penricei Rothschild 1895 Penric's waterbuck
LR/cd V
+
+
+
?
Kobus leche Gray 1850 Red lechwe
LR/cd V
+
+
+
+
Kobus kafuensis Haltenorth 1964 Kafue lechwe
VU D2 Ex
Kobus vardoni (Livingstone 1857) Puku
LR/cd V
E?
+
+
+
E
Hippotragus equinus (Desmarest 1804) Roan antelope
LR/nt V
+
+
+
+
+
Hippotragus niger (Harris 1838) Sable antelope
LR/nt
+
+
+
+
+
Oryx gazella (Linnaeus 1758) Gemsbok
LR/cd
Tribe Reduncini
+
+
+
Tribe Hippotragini
+
+
Tribe Alcelaphini
Connachaetes taurinus (Burchell 1823) Blue wildebeest
LR/cd
+
+
+
+
+
Damaliscus lunatus (Burchell 1823) Tsessebe
LR/nt V
+
+
+
+
+
Alcelaphus cama (Pallas 1776) Red hartebeest
LR/cd
+
+
+
Sigmoceros lichtensteini (Peters 1849) Lichtenstein's hartebeest
LR/cd
+
+
+
Tribe Antilopini
Antidorcas marsupialis (Zimmerman 1780) Springbok
LR/nt
+
Tribe Aepyceroni
Aepyceros melampus (Lichtenstein 1812) Impala
LR/nt
+
+
+
+
+
+
+
Tribe Neotragini
Oreotragus oreotragus (Zimmerman 1783) Klipspringer
LR/cd V
+
Ourebia ourebi (Zimmerman 1783) Oribi
V
+
+
+
+
+
Raphicerus campestris (Thunberg 1811) Stenbuck
LR/nt
+
+
+
+
+
Raphicerus sharpei Thomas 1897 Sharpe's grysbuck
V
+
+
+
+
+
Order RODENTIA
Thryonomys swinderianus (Temminck 1827) Greater canerat
LR/nt
Thryonomys gregorianus (Thomas 1894) Lesser canerat
DD
+
+
+
+
Family Bathyergidae
Cryptomys damarensis (Ogilby 1838) Damara molerat
LR/nt
+
+
+
+
+
265
Taxon
Conserv.
status
Cryptomys kafuensis Burda et al. 1999 Kafue molerat
DD
+
Cryptomys sp. molerat (Livingstone environs)
DD
+
Ang
Bot
Nam Zam Zim
Family Hystricidae
Hystrix africaeaustralis Peters 1852 Porcupine
LR/nt
+
+
+
+
+
Family Sciuridae (squirrels)
Anomalurus derbianus (Gray 1842) Scaly-tailed flying squirrel
LR/nt V
+
Heliosciurus gambianus (Ogilby 1835) Gambian sun squirrel
LR/nt V
+
Paraxerus cepapi (A.Smith 1836) Tree squirrel
LR/nt
Xerus inaurus (Zimmermann 1780) Ground squirrel
LR/nt
+
+
+
+
+
+
Family Peditidae
Pedetes capensis (Forster 1778) Springhare
LR/nt
+
+
+
+
+
Graphiurus murinus (Desmarest 1822) Woodland dormouse
LR/nt
+
+
+
+
+
Graphiurus platyops Thomas 1897 Rock dormouse
LR/nt
+
+
+
+
+
+
+
+
+
+
Family Graphiuridae (dormice)
Family Muridae (rats and mice)
Saccostomys campestris Peters 1846 Pouched mouse
LR/nt
+
+
Dendromus mystacalis Heughlin 1863 Lesser climbing mouse
LR/nt
Dendromus melanotis A.Smith 1834 Grey climbing mouse
LR/nt
+
+
+
Dendromus mesomelas (Brants 1827) Chestnut climbing mouse
LR/nt
+
+
+
Dendromus nyikae Wroughton 1909 Climbing mouse
LR/nt
Steatomys krebsi Peters 1852 Kreb's fat mouse
LR/nt
+
Steatomys pratensis Peters 1846 Fat mouse
LR/nt
Steatomys parvus Rhoads 1896 Tiny fat mouse
LR/nt
Otomys angoniensis Wroughton 1906 Angoni vlei rat
LR/nt
Otomys maximus Roberts 1924 Large vlei rat
LR/nt
+
Desmodillus auricularis (A.Smith 1834) Namaqua gerbil
LR/nt
+
Gerbillurus paeba (A.Smith 1836) Hairy-footed gerbil
LR/nt
+
Tatera valida (Bocage 1890) Bocage's gerbil
LR/nt
+
Tatera brantsi (A.Smith 1836) Brants' gerbil
LR/nt
+
+
+
+
+
Tatera leucogaster (Peters 1852) Bushveld gerbil
LR/nt
+
+
+
+
+
Mus musculus Linnaeus 1758 House mouse (introduced)
LR/nt
+
+
Mus minutoides A.Smith 1834 Pygmy mouse
LR/nt
+
+
+
+
+
Mus indutus (Thomas 1910) Desert Pygmy mouse
LR/nt
+
+
+
+
+
Mastomys coucha (A.Smith 1836) Multimammate mouse
LR/nt
+
+
+
+
+
Mastomys natalensis (A.Smith 1834) Natal multimammate mouse
LR/nt
+
+
+
+
+
Mastomys shortridgei (St Leger 1933) Shortridge's mouse
LR/nt
+
+
Rattus rattus (Linnaeus 1758) House rat (introduced)
LR/nt
+
+
+
+
+
Thallomys paedulcus (Sundevall 1846) Tree rat
LR/nt
+
+
+
+
+
Dasymys incomtus (Sundevall 1847) Water rat
DD
+
+
+
+
+
Aethomys namaquensis (A.Smith 1834) Namaqua rock rat
LR/nt
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
266
Taxon
Conserv.
status
Aethomys chrysophilus (De Winton 1897) Red veld rat
LR/nt
+
+
+
+
Pellomys fallax (Peters 1852) Groove-toothed rat
LR/nt
+
+
+
+
Lemniscomys rosalia (Thomas 1904) Single-striped mouse
LR/nt
+
+
+
+
Arvicanthus niloticus (Desmarest 1822)
LR/nt
Rhabdomys pumilio (Sparrmann 1784) Striped mouse
DD
Zelotomys hildegardeae (Thomas 1902) Hildegarde's rat
LR/nt
Zelatomys woosnami (Schwann 1906) Woosnam's desert rat
LR/nt
Grammomys dolichurus (Smuts 1832) Woodland rat
LR/nt
Acomys spinosissimus (Peters 1852) Spiny mouse
LR/nt
Ang Bot
+
Nam Zam Zim
+
+
+
+
+
+
+
+
+
+
Order PHOLIDOTA (pangolins)
Manis temminckii Smuts 1832 Pangolin
LR/nt V
+
+
+
+
+
Lepus saxatalis F.Cuvier 1823 Scrub hare
LR/nt
+
+
+
+
+
Lepus capensis Linnaeus 1758 Cape Hare
LR/nt
Pronolagus randensis Jameson 1907 Jameson's red rock rabbit
LR/nt
LEPORIDAE (hares and rabbits)
+
+
MACROSCELIDAE (elephant shrews)
Petrodomus tetradactylus Peters 1846 Four-toed elephant shrew
LR/nt
Elephantulus brachyrhynchus (A.Smith 1836) Short-snouted
elephant shrew
LR/nt
Elephantulus intufti (A.Smith 1836) Bushveld elephant shrew
LR/nt
Elephantulus myurus Thomas & Schwann 1906 Rock elephant
shrew
LR/nt
TOTAL = 197 species
+
+
+
+
+
+
+
91
149
118
162
150

Mammals - The Zambezi Society

Transcription

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